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||
<span class="search-input-link user-guide-link" role="navigation" aria-label="User guide">
|
||
<a href="/help/"
|
||
data-ga-category="featured_link"
|
||
data-ga-action="user_guide">
|
||
User Guide
|
||
</a>
|
||
</span>
|
||
|
||
</div>
|
||
</div>
|
||
|
||
|
||
</form>
|
||
|
||
|
||
<div id="article-page-header">
|
||
<div id="article-top-actions-bar" class="article-top-actions-bar">
|
||
<div class="inner-wrap">
|
||
<div class="actions-container">
|
||
<div class="return-to-search">
|
||
|
||
|
||
</div>
|
||
|
||
<div class="multiple-results-actions " role="region" aria-label="save, email, send to">
|
||
<button id="save-results-panel-trigger"
|
||
type="button"
|
||
class="save-results save-results-panel-trigger"
|
||
aria-expanded="false"
|
||
aria-controls="save-action-panel"
|
||
data-ga-category="save_share"
|
||
data-ga-action="save"
|
||
data-ga-label="open">
|
||
Save
|
||
</button><button id="email-results-panel-trigger"
|
||
type="button"
|
||
class="email-results email-results-panel-trigger"
|
||
aria-expanded="false"
|
||
aria-controls="email-action-panel"
|
||
data-ga-category="save_share"
|
||
data-ga-action="email"
|
||
data-ga-label="open">
|
||
Email
|
||
</button><div class="more-actions dropdown-block"><button id="more-actions-trigger"
|
||
type="button"
|
||
class="trigger more-actions-trigger"
|
||
ref="linksrc=show_moreactions_btn"
|
||
aria-label="Send to"
|
||
aria-controls="more-actions-dropdown">Send to</button><div id="more-actions-dropdown" class="dropdown dropdown-container" hidden><div class="content"><ul class="more-actions-links"><li><a class="clipboard-panel-trigger dropdown-block-link "
|
||
href="#"
|
||
role="button"
|
||
data-ga-category="save_share"
|
||
data-ga-action="clipboard"
|
||
data-ga-label="send">
|
||
Clipboard
|
||
</a></li><li><a role="button" class="dropdown-block-link bibliography-trigger-target" href="https://account.ncbi.nlm.nih.gov/?back_url=https%3A%2F%2Fpubmed.ncbi.nlm.nih.gov%2F23028350%2F%23open-bibliography-panel">My Bibliography</a></li><li><a role="button" class="dropdown-block-link collections-trigger-target" href="https://account.ncbi.nlm.nih.gov/?back_url=https%3A%2F%2Fpubmed.ncbi.nlm.nih.gov%2F23028350%2F%23open-collections-panel">Collections</a></li><li><a role="button" class="citation-manager-panel-trigger dropdown-block-link citation-manager-trigger-target" href="#">Citation manager</a></li></ul></div></div></div>
|
||
</div>
|
||
|
||
|
||
|
||
<div class="display-options" data-format-key="format"
|
||
data-default-format-value="abstract"
|
||
data-current-format-value="abstract" role="region">
|
||
<button
|
||
class="trigger usa-button"
|
||
title="Change format"
|
||
data-ga-category="display_options"
|
||
data-ga-action="display_options_dialog">
|
||
<span class="button-label">Display options</span>
|
||
</button>
|
||
|
||
<div class="dropdown" hidden>
|
||
<div class="title">
|
||
Display options
|
||
</div>
|
||
|
||
<div class="content">
|
||
|
||
<div class="format-container">
|
||
<span class="option-label">
|
||
Format
|
||
</span>
|
||
<select name="format" data-initial-value="abstract" aria-label="Display format" id="article-display-format">
|
||
<option value="abstract">Abstract</option>
|
||
|
||
<option value="pubmed">PubMed</option>
|
||
|
||
<option value="pmid">PMID</option>
|
||
|
||
</select>
|
||
</div>
|
||
|
||
</div>
|
||
</div>
|
||
</div>
|
||
<input type="hidden" name="csrfmiddlewaretoken" value="vzMy7zZkDk3vgebaPSut8k1jcJTdfjRPMmwDKpsGdVQZaFh0sb7vCsxXKNBp8Up9">
|
||
</div>
|
||
</div>
|
||
</div>
|
||
|
||
<div id="article-panels-container" class="article-panels-container">
|
||
<div id="save-action-panel" class="save-action-panel action-panel" aria-hidden="true">
|
||
<div class="inner-wrap">
|
||
<h2 class="action-panel-heading">
|
||
Save citation to file
|
||
</h2>
|
||
<form id="save-action-panel-form"
|
||
class="action-panel-content action-form"
|
||
action="/results-export-ids/"
|
||
data-by-search-action="/results-export-search-data/"
|
||
data-by-ids-action="/results-export-ids/"
|
||
method="post"
|
||
data-by-search-method="post"
|
||
data-by-ids-method="post">
|
||
|
||
<input type="hidden" name="csrfmiddlewaretoken" value="vzMy7zZkDk3vgebaPSut8k1jcJTdfjRPMmwDKpsGdVQZaFh0sb7vCsxXKNBp8Up9">
|
||
|
||
|
||
|
||
<div class="action-panel-control-wrap">
|
||
<label for="save-action-format" class="action-panel-label">
|
||
Format:
|
||
</label>
|
||
<select id="save-action-format" name="results-format" class="action-panel-selector">
|
||
<option value="summary-text">Summary (text)</option>
|
||
<option value="pubmed-text">PubMed</option>
|
||
<option value="pmid">PMID</option>
|
||
<option value="abstract">Abstract (text)</option>
|
||
<option value="csv">CSV</option>
|
||
</select>
|
||
</div>
|
||
|
||
<div class="action-panel-actions">
|
||
<button class="action-panel-submit"
|
||
type="submit"
|
||
data-loading-label="Saving..."
|
||
data-ga-category="save_share"
|
||
data-ga-action="save"
|
||
data-ga-label="save">
|
||
Create file
|
||
</button>
|
||
<button class="action-panel-cancel"
|
||
aria-label="Close 'Save citations to file' panel"
|
||
ref="linksrc=close_save_panel"
|
||
aria-controls="save-action-panel"
|
||
aria-expanded="false"
|
||
data-ga-category="save_share"
|
||
data-ga-action="save"
|
||
data-ga-label="cancel">
|
||
Cancel
|
||
</button>
|
||
</div>
|
||
</form>
|
||
</div>
|
||
</div>
|
||
|
||
|
||
|
||
|
||
<div id="email-action-panel"
|
||
class="email-action-panel action-panel"
|
||
aria-hidden="true"
|
||
data-recaptcha-script-url="https://www.recaptcha.net/recaptcha/enterprise.js?render=explicit&onload=onloadCaptchaCallback"
|
||
data-recaptcha-enabled="true">
|
||
<div class="inner-wrap">
|
||
<h2 class="action-panel-heading">
|
||
Email citation
|
||
</h2>
|
||
<form id="email-action-panel-form"
|
||
class="action-panel-content action-form"
|
||
action="/send-email/"
|
||
data-by-search-action="/results-export-email-by-search-data/"
|
||
data-by-ids-action="/send-email/"
|
||
data-by-search-method="post"
|
||
data-by-ids-method="post"
|
||
method="post"
|
||
>
|
||
<input type="hidden" name="csrfmiddlewaretoken" value="vzMy7zZkDk3vgebaPSut8k1jcJTdfjRPMmwDKpsGdVQZaFh0sb7vCsxXKNBp8Up9">
|
||
|
||
<div class="action-panel-control-wrap">
|
||
<span class="action-panel-label">
|
||
Subject:
|
||
</span>
|
||
<span class="email-subject" id="email-subject" aria-label="Email subject">1 selected item: 23028350 - PubMed</span>
|
||
<input type="hidden" name="email-subject" value="1 selected item: 23028350"/>
|
||
<input type="hidden" name="email-subject-hash" value="136cd3ee04ceabf4fbe75ec5fa82348bcdc97b2bb108ddc7235e22718573e379"/>
|
||
</div>
|
||
|
||
<div class="action-panel-control-wrap">
|
||
<label for="email-to" class="action-panel-label required-field-asterisk">
|
||
To:
|
||
</label>
|
||
<input type="email" aria-label="Recipient Email Address" placeholder="email@example.com" id="email-to" class="email-to" pattern="^[a-zA-Z0-9_.+-]+@[a-zA-Z0-9-]+\.[a-zA-Z0-9-.]+$" maxlength="100" required>
|
||
</div>
|
||
|
||
<div class="action-panel-control-wrap">
|
||
<label for="email-from" class="action-panel-label">
|
||
From:
|
||
</label>
|
||
<input type="email" aria-label="Sender Email Address" placeholder="email@example.com" id="email-from" class="email-from" pattern="^[a-zA-Z0-9_.+-]+@[a-zA-Z0-9-]+\.[a-zA-Z0-9-.]+$" maxlength="256">
|
||
</div>
|
||
|
||
|
||
|
||
<div class="action-panel-control-wrap">
|
||
<label for="email-citation-format" class="action-panel-label">
|
||
Format:
|
||
</label>
|
||
<select id="email-citation-format" name="citation-format" class="action-panel-selector email-citation-format">
|
||
<option selected="selected" value="summary">Summary</option>
|
||
<option value="summary-text">Summary (text)</option>
|
||
<option value="abstract">Abstract</option>
|
||
<option value="abstract-text">Abstract (text)</option>
|
||
|
||
</select>
|
||
</div>
|
||
<div class="include-supplemental-container">
|
||
<input type="checkbox" aria-label="Include MeSH and other data" name="include-supplemental" id="email-include-supplemental" class="email-include-supplemental">
|
||
<label for="email-include-supplemental" class="email-include-supplemental-label">MeSH and other data</label>
|
||
</div>
|
||
|
||
|
||
|
||
<div class="form-field recaptcha ">
|
||
|
||
|
||
|
||
<div class="g-recaptcha" id="id-recaptcha" data-sitekey="6LfsWHMdAAAAAClKbtOpjQ2pMjgsGxvv7NdZW9uI"></div>
|
||
|
||
|
||
|
||
|
||
|
||
</div>
|
||
<div id="captcha-error-message" class="usa-input-error-message captcha-validation-message" role="alert"></div>
|
||
|
||
|
||
|
||
<div class="action-panel-actions">
|
||
<button class="action-panel-submit"
|
||
type="submit"
|
||
data-loading-label="Sending..."
|
||
data-ga-category="save_share"
|
||
data-ga-action="email"
|
||
data-ga-label="send">
|
||
Send email
|
||
</button>
|
||
<button class="action-panel-cancel"
|
||
aria-label="Close 'Email citations' panel"
|
||
ref="linksrc=close_email_panel"
|
||
aria-controls="email-action-panel"
|
||
aria-expanded="false"
|
||
data-ga-category="save_share"
|
||
data-ga-action="email"
|
||
data-ga-label="cancel">
|
||
Cancel
|
||
</button>
|
||
</div>
|
||
<input type="hidden" name="email-search-details" value="" />
|
||
<input type="hidden" name="email-search-details-hash" value="0e42663a6c3bd85498fcb88798998fed7bfdc45d457db35281e41afe13cc0524" />
|
||
</form>
|
||
</div>
|
||
</div>
|
||
|
||
<div id="collections-action-panel"
|
||
class="collections-action-panel action-panel in-progress-dots-panel"
|
||
aria-hidden="true"
|
||
data-collections-open-panel-enabled="false"
|
||
data-collections-open-panel-url-hash="#open-collections-panel">
|
||
<div class="inner-wrap">
|
||
<h3 class="action-panel-heading">
|
||
Add to Collections
|
||
</h3>
|
||
|
||
|
||
<form id="collections-action-panel-form"
|
||
class="collections-action-panel-form action-panel-content action-form action-panel-smaller-selectors"
|
||
data-existing-collections-url="/list-existing-collections/"
|
||
data-add-to-existing-collection-url="/add-to-existing-collection/"
|
||
data-create-and-add-to-new-collection-url="/create-and-add-to-new-collection/"
|
||
data-get-article-ids-by-search-url="/get-article-ids-by-search/"
|
||
data-myncbi-max-collection-name-length="100"
|
||
data-add-to-collection-max-amount="1000"
|
||
data-collections-root-url="https://www.ncbi.nlm.nih.gov/myncbi/collections/">
|
||
|
||
<input type="hidden" name="csrfmiddlewaretoken" value="vzMy7zZkDk3vgebaPSut8k1jcJTdfjRPMmwDKpsGdVQZaFh0sb7vCsxXKNBp8Up9">
|
||
|
||
|
||
|
||
<div class="choice-group" role="radiogroup">
|
||
<ul class="radio-group-items">
|
||
<li>
|
||
<input type="radio"
|
||
id="collections-action-panel-new"
|
||
class="collections-new"
|
||
name="collections"
|
||
value="new"
|
||
data-ga-category="save_share"
|
||
data-ga-action="collections"
|
||
data-ga-label="collections_radio_new">
|
||
<label for="collections-action-panel-new">Create a new collection</label>
|
||
</li>
|
||
<li>
|
||
<input type="radio"
|
||
id="collections-action-panel-existing"
|
||
class="collections-existing"
|
||
name="collections"
|
||
value="existing"
|
||
checked="true"
|
||
data-ga-category="save_share"
|
||
data-ga-action="collections"
|
||
data-ga-label="collections_radio_existing">
|
||
<label for="collections-action-panel-existing">Add to an existing collection</label>
|
||
</li>
|
||
</ul>
|
||
</div>
|
||
|
||
<div class="controls-wrapper">
|
||
<div class="action-panel-control-wrap new-collections-controls">
|
||
<label for="collections-action-panel-add-to-new" class="action-panel-label required-field-asterisk">
|
||
Name your collection:
|
||
</label>
|
||
<input
|
||
type="text"
|
||
name="add-to-new-collection"
|
||
id="collections-action-panel-add-to-new"
|
||
class="collections-action-add-to-new"
|
||
pattern="[^"&=<>\/]*" title="The following characters are not allowed in the Name field: "&=<>/"
|
||
maxlength="100"
|
||
data-ga-category="save_share"
|
||
data-ga-action="create_collection"
|
||
data-ga-label="non_favorties_collection">
|
||
<div class="collections-new-name-too-long usa-input-error-message selection-validation-message">
|
||
Name must be less than 100 characters
|
||
</div>
|
||
</div>
|
||
<div class="action-panel-control-wrap existing-collections-controls">
|
||
<label for="collections-action-panel-add-to-existing" class="action-panel-label">
|
||
Choose a collection:
|
||
</label>
|
||
<select id="collections-action-panel-add-to-existing"
|
||
class="action-panel-selector collections-action-add-to-existing"
|
||
data-ga-category="save_share"
|
||
data-ga-action="select_collection"
|
||
data-ga-label="($('#collections-action-add-to-existing').val() === 'Favorites') ? 'Favorites' : 'non_favorites_collection'">
|
||
</select>
|
||
<div class="collections-retry-load-on-error usa-input-error-message selection-validation-message">
|
||
Unable to load your collection due to an error<br>
|
||
<a href="#">Please try again</a>
|
||
</div>
|
||
</div>
|
||
</div>
|
||
|
||
<div class="action-panel-actions">
|
||
<button class="action-panel-submit"
|
||
type="submit"
|
||
data-loading-label="Adding..."
|
||
data-pinger-ignore
|
||
data-ga-category="save_share"
|
||
data-ga-action="collections"
|
||
data-ga-label="add">
|
||
Add
|
||
</button>
|
||
<button class="action-panel-cancel"
|
||
aria-label="Close 'Add to Collections' panel"
|
||
ref="linksrc=close_collections_panel"
|
||
aria-controls="collections-action-panel"
|
||
aria-expanded="false"
|
||
data-ga-category="save_share"
|
||
data-ga-action="collections"
|
||
data-ga-label="cancel">
|
||
Cancel
|
||
</button>
|
||
</div>
|
||
</form>
|
||
</div>
|
||
</div>
|
||
|
||
<div id="bibliography-action-panel"
|
||
class="bibliography-action-panel action-panel in-progress-dots-panel"
|
||
aria-hidden="true"
|
||
data-bibliography-open-panel-enabled="false"
|
||
data-bibliography-open-panel-url-hash="#open-bibliography-panel">
|
||
<div class="inner-wrap">
|
||
<h3 class="action-panel-heading">
|
||
Add to My Bibliography
|
||
</h3>
|
||
<form id="bibliography-action-panel-form"
|
||
class="bibliography-action-panel-form action-panel-content action-form action-panel-smaller-selectors"
|
||
data-add-to-bibliography-max-amount="100"
|
||
data-add-to-bibliography-batch-size="10"
|
||
data-bibliography-delegates-url="/list-bibliography-delegates/"
|
||
data-add-to-bibliography-url="/add-to-bibliography/"
|
||
data-get-article-ids-by-search-url="/get-article-ids-by-search/"
|
||
data-mybib-root-url="https://www.ncbi.nlm.nih.gov/myncbi/collections/mybibliography/">
|
||
|
||
<input type="hidden" name="csrfmiddlewaretoken" value="vzMy7zZkDk3vgebaPSut8k1jcJTdfjRPMmwDKpsGdVQZaFh0sb7vCsxXKNBp8Up9">
|
||
|
||
|
||
|
||
<div class="action-panel-control-wrap bibliographies-controls">
|
||
<div class="choice-group">
|
||
<ul class="bibliographies-action-add radio-group-items">
|
||
<li>
|
||
<input name="bibliography" id="my-bibliography" class="my-bibliography" type="radio" checked/>
|
||
<label for="my-bibliography">My Bibliography</label>
|
||
</li>
|
||
</ul>
|
||
</div>
|
||
</div>
|
||
|
||
<div class="bibliographies-retry-load-on-error usa-input-error-message selection-validation-message">
|
||
Unable to load your delegates due to an error<br>
|
||
<a href="#">Please try again</a>
|
||
</div>
|
||
|
||
<div class="action-panel-actions">
|
||
<button class="action-panel-submit"
|
||
type="submit"
|
||
data-loading-label="Adding..."
|
||
data-pinger-ignore>
|
||
Add
|
||
</button>
|
||
<button class="action-panel-cancel"
|
||
aria-label="Close 'Add to bibliography' panel"
|
||
ref="linksrc=close_bibliography_panel"
|
||
aria-controls="bibliography-action-panel"
|
||
aria-expanded="false"
|
||
data-ga-category="save_share"
|
||
data-ga-action="mybib"
|
||
data-ga-label="cancel">
|
||
Cancel
|
||
</button>
|
||
</div>
|
||
</form>
|
||
</div>
|
||
</div>
|
||
|
||
|
||
|
||
|
||
<div id="saved-search-action-panel" class="saved-search-action-panel action-panel " aria-hidden="true"
|
||
data-saved-search-open-panel-enabled="false"
|
||
data-saved-search-open-panel-url-hash="#open-saved-search-panel">
|
||
<div class="inner-wrap">
|
||
<h2 class="action-panel-heading">
|
||
Your saved search
|
||
</h2>
|
||
|
||
<form id="saved-search-action-panel-form"
|
||
class="saved-search-action-panel-form action-panel-content action-form"
|
||
data-create-saved-search-url="/create-saved-search/"
|
||
data-try-search-terms-url="/try-search-term/"
|
||
data-saved-search-root-url="https://www.ncbi.nlm.nih.gov/myncbi/searches/">
|
||
|
||
<input type="hidden" name="csrfmiddlewaretoken" value="vzMy7zZkDk3vgebaPSut8k1jcJTdfjRPMmwDKpsGdVQZaFh0sb7vCsxXKNBp8Up9">
|
||
|
||
<div class="action-panel-control-wrap">
|
||
<label for="saved-search-name" class="action-panel-label saved-search-name-label required-field-asterisk">
|
||
Name of saved search:
|
||
</label>
|
||
<input maxlength="200"
|
||
type="text"
|
||
name="saved-search-name"
|
||
id="saved-search-name"
|
||
class="saved-search-name"
|
||
value=""
|
||
required
|
||
pattern="[^"&=<>\/]*" title="The following characters are not allowed in the Name field: "&=<>/">
|
||
</div>
|
||
|
||
<div class="action-panel-control-wrap">
|
||
<label for="saved-search-term" class="action-panel-label required-field-asterisk">
|
||
Search terms:
|
||
</label>
|
||
<textarea name="saved-search-term" id="saved-search-term" class="saved-search-term" required></textarea>
|
||
</div>
|
||
<div class="test-search-term-wrap">
|
||
<a href="#" class="try-search-term">Test search terms</a>
|
||
</div>
|
||
<div class="choice-group action-panel-extra-margin-top">
|
||
<span class="action-panel-label" id="fieldset-label">
|
||
Would you like email updates of new search results?
|
||
</span>
|
||
<fieldset id="saved-search-alert" aria-describedby="fieldset-label">
|
||
<legend class="usa-sr-only">Saved Search Alert Radio Buttons</legend>
|
||
<ul class="radio-group-items">
|
||
<li>
|
||
<input type="radio" id="saved-search-alert-yes" class="saved-search-alert-yes" name="saved-search-alert" value="yes" checked>
|
||
<label for="saved-search-alert-yes" class="action-panel-label">Yes</label>
|
||
</li>
|
||
<li>
|
||
<input aria-label="No radio input" type="radio" id="saved-search-alert-no" class="saved-search-alert-no" name="saved-search-alert" value="no">
|
||
<label for="saved-search-alert-no" class="action-panel-label">No</label>
|
||
</li>
|
||
</ul>
|
||
</fieldset>
|
||
</div>
|
||
|
||
<div class="alert-schedule-wrap">
|
||
<div class="action-panel-control-wrap">
|
||
<label class="action-panel-label">
|
||
Email:
|
||
</label>
|
||
<span aria-label="Email address" id="saved-search-email" class="action-panel-label"><span class="action-panel-label-bold"></span> (<a class="myncbi-account-settings" href="https://www.ncbi.nlm.nih.gov/account/settings/">change</a>)</span>
|
||
</div>
|
||
<div class="action-panel-control-wrap action-panel-extra-margin-top">
|
||
<label for="saved-search-frequency" class="action-panel-label">
|
||
Frequency:
|
||
</label>
|
||
<select id="saved-search-frequency" class="no-border-panel-selector saved-search-frequency">
|
||
<option value="monthly">Monthly</option>
|
||
<option value="weekly">Weekly</option>
|
||
<option value="daily">Daily</option>
|
||
</select>
|
||
</div>
|
||
<div class="action-panel-control-wrap saved-search-monthly-additional">
|
||
<label for="saved-search-monthly-on-day" class="action-panel-label">
|
||
Which day?
|
||
</label>
|
||
<select id="saved-search-monthly-on-day" class="no-border-panel-selector">
|
||
<option value="Sunday">The first Sunday</option>
|
||
<option value="Monday">The first Monday</option>
|
||
<option value="Tuesday">The first Tuesday</option>
|
||
<option value="Wednesday">The first Wednesday</option>
|
||
<option value="Thursday">The first Thursday</option>
|
||
<option value="Friday">The first Friday</option>
|
||
<option value="Saturday">The first Saturday</option>
|
||
<option value="day">The first day</option>
|
||
<option value="weekday">The first weekday</option>
|
||
</select>
|
||
</div>
|
||
<div class="action-panel-control-wrap saved-search-weekly-additional">
|
||
<label for="saved-search-weekly-on-day" class="action-panel-label">
|
||
Which day?
|
||
</label>
|
||
<select id="saved-search-weekly-on-day" class="no-border-panel-selector saved-search-weekly-on-day">
|
||
<option value="Sunday">Sunday</option>
|
||
<option value="Monday">Monday</option>
|
||
<option value="Tuesday">Tuesday</option>
|
||
<option value="Wednesday">Wednesday</option>
|
||
<option value="Thursday">Thursday</option>
|
||
<option value="Friday">Friday</option>
|
||
<option value="Saturday">Saturday</option>
|
||
</select>
|
||
</div>
|
||
<div class="action-panel-control-wrap">
|
||
<label for="saved-search-report" class="action-panel-label">
|
||
Report format:
|
||
</label>
|
||
<select id="saved-search-report" class="no-border-panel-selector saved-search-report">
|
||
<option value="DocSum">Summary</option>
|
||
<option value="DocSumText">Summary (text)</option>
|
||
<option value="Abstract">Abstract</option>
|
||
<option value="AbstractText">Abstract (text)</option>
|
||
<option value="MEDLINE">PubMed</option>
|
||
</select>
|
||
</div>
|
||
<div class="action-panel-control-wrap">
|
||
<label for="saved-search-amount" class="action-panel-label">
|
||
Send at most:
|
||
</label>
|
||
<select id="saved-search-amount" class="no-border-panel-selector saved-search-amount">
|
||
<option value="1">1 item</option>
|
||
<option value="5" selected>5 items</option>
|
||
<option value="10">10 items</option>
|
||
<option value="20">20 items</option>
|
||
<option value="50">50 items</option>
|
||
<option value="100">100 items</option>
|
||
<option value="200">200 items</option>
|
||
</select>
|
||
</div>
|
||
<div>
|
||
<input type="checkbox" id="saved-search-send-if-no-result" class="saved-search-send-if-no-result" name="saved-search-send-if-no-result">
|
||
<label for="saved-search-send-if-no-result" class="action-panel-label smaller-checkbox">
|
||
Send even when there aren't any new results
|
||
</label>
|
||
</div>
|
||
<div class="action-panel-control-wrap option-text-in-email-wrap">
|
||
<label for="saved-search-email-text" class="action-panel-label">
|
||
Optional text in email:
|
||
</label>
|
||
<textarea name="saved-search-email-text"
|
||
id="saved-search-email-text"
|
||
class="saved-search-email-text"></textarea>
|
||
</div>
|
||
|
||
</div>
|
||
|
||
<div class="action-panel-actions">
|
||
<button class="action-panel-submit"
|
||
type="submit"
|
||
data-loading-label="Saving..."
|
||
data-ga-category="save_share"
|
||
data-ga-action="alert"
|
||
data-ga-label="save">
|
||
Save
|
||
</button>
|
||
<button class="action-panel-cancel"
|
||
aria-label="Close 'Your saved search' panel"
|
||
ref="linksrc=close_saved_search_panel"
|
||
aria-controls="saved-search-action-panel"
|
||
aria-expanded="false"
|
||
data-ga-category="save_share"
|
||
data-ga-action="alert"
|
||
data-ga-label="cancel">
|
||
Cancel
|
||
</button>
|
||
</div>
|
||
</form>
|
||
</div>
|
||
</div>
|
||
|
||
<div id="citation-manager-action-panel" class="citation-manager-action-panel action-panel" aria-hidden="true">
|
||
<div class="inner-wrap">
|
||
<h2 class="action-panel-heading">
|
||
Create a file for external citation management software
|
||
</h2>
|
||
<form id="citation-manager-action-panel-form"
|
||
class="action-panel-content action-form"
|
||
action="/results-export-ids/"
|
||
data-by-search-action="/results-export-search-data/"
|
||
data-by-ids-action="/results-export-ids/"
|
||
method="post"
|
||
data-by-search-method="post"
|
||
data-by-ids-method="post">
|
||
|
||
<input type="hidden" name="csrfmiddlewaretoken" value="vzMy7zZkDk3vgebaPSut8k1jcJTdfjRPMmwDKpsGdVQZaFh0sb7vCsxXKNBp8Up9">
|
||
|
||
|
||
|
||
<input name="results-format" type="hidden" value="pubmed"/>
|
||
|
||
<div class="action-panel-actions">
|
||
<button class="action-panel-submit"
|
||
type="submit"
|
||
data-loading-label="Sending..."
|
||
data-ga-category="save_share"
|
||
data-ga-action="citation_manager"
|
||
data-ga-label="save">
|
||
Create file
|
||
</button>
|
||
<button class="action-panel-cancel"
|
||
aria-label="Close 'Send citations to citation manager' panel"
|
||
ref="linksrc=close_citation_manager_panel"
|
||
aria-controls="citation-manager-action-panel"
|
||
aria-expanded="false"
|
||
data-ga-category="save_share"
|
||
data-ga-action="citation_manager"
|
||
data-ga-label="cancel">
|
||
Cancel
|
||
</button>
|
||
</div>
|
||
</form>
|
||
</div>
|
||
</div>
|
||
|
||
|
||
|
||
|
||
<div id="rss-action-panel" class="rss-action-panel action-panel " aria-hidden="true">
|
||
<div class="inner-wrap">
|
||
<h2 class="action-panel-heading">
|
||
Your RSS Feed
|
||
</h2>
|
||
|
||
<form id="rss-action-panel-form"
|
||
class="rss-action-panel-form action-panel-content action-form"
|
||
data-create-rss-feed-url="/create-rss-feed-url/"
|
||
data-search-form-term-value="">
|
||
|
||
<input type="hidden" name="csrfmiddlewaretoken" value="vzMy7zZkDk3vgebaPSut8k1jcJTdfjRPMmwDKpsGdVQZaFh0sb7vCsxXKNBp8Up9">
|
||
|
||
<div class="action-panel-control-wrap">
|
||
<label for="rss-name" class="action-panel-label required-field-asterisk">
|
||
Name of RSS Feed:
|
||
</label>
|
||
<input maxlength="200"
|
||
placeholder="Name"
|
||
type="text"
|
||
name="rss-name"
|
||
id="rss-name"
|
||
class="rss-name"
|
||
value=''
|
||
required
|
||
pattern="[^"&=<>\/]*" title="The following characters are not allowed in the Name field: "&=<>/">
|
||
</div>
|
||
|
||
<div class="rss-limit-wrap">
|
||
<div class="action-panel-control-wrap action-panel-extra-margin-top">
|
||
<label for="rss-limit" class="action-panel-label">
|
||
Number of items displayed:
|
||
</label>
|
||
<select id="rss-limit" class="no-border-panel-selector rss-limit">
|
||
<option value="5">5</option>
|
||
<option value="10">10</option>
|
||
<option value="15" selected="selected">15</option>
|
||
<option value="20">20</option>
|
||
<option value="50">50</option>
|
||
<option value="100">100</option>
|
||
</select>
|
||
</div>
|
||
</div>
|
||
|
||
<div class="action-panel-actions">
|
||
<button class="action-panel-submit"
|
||
type="submit"
|
||
data-loading-label="Creating..."
|
||
data-ga-category="save_share"
|
||
data-ga-action="alert"
|
||
data-ga-label="save">
|
||
Create RSS
|
||
</button>
|
||
<button class="action-panel-cancel"
|
||
aria-label="Close 'Your RSS' panel"
|
||
ref="linksrc=close_rss_panel"
|
||
aria-controls="rss-action-panel"
|
||
aria-expanded="false"
|
||
data-ga-category="save_share"
|
||
data-ga-action="alert"
|
||
data-ga-label="cancel">
|
||
Cancel
|
||
</button>
|
||
</div>
|
||
|
||
<div class="action-panel-control-wrap rss-link-copy-wrap">
|
||
<label for="rss-link" class="usa-sr-only">RSS Link</label>
|
||
<input placeholder="Your RSS Feed Link" type="text" name="rss-link" id="rss-link" class="rss-link" title="RSS Link">
|
||
<button
|
||
type="button"
|
||
disabled
|
||
class="rss-link-copy-button disabled"
|
||
data-ga-category="save_share"
|
||
data-ga-action="rss"
|
||
data-ga-label="copy">
|
||
Copy
|
||
</button>
|
||
</div>
|
||
</form>
|
||
</div>
|
||
</div>
|
||
|
||
</div>
|
||
</div>
|
||
|
||
<div class="article-page" id="article-page" data-article-pmid="23028350">
|
||
<aside class="page-sidebar">
|
||
<div class="inner-wrap">
|
||
|
||
<div class="full-text-links">
|
||
<div class="full-view">
|
||
<h3 class="title">
|
||
Full text links
|
||
</h3>
|
||
<div class="full-text-links-list">
|
||
<a class="link-item
|
||
|
||
dialog-focus"
|
||
href="https://dx.plos.org/10.1371/journal.pgen.1002938"
|
||
target="_blank"
|
||
rel="noopener"
|
||
|
||
ref="linksrc=fulltextorjournal_fulltext&is_pmc=False&PrId=4656&itool=Abstract-def&log$=linkouticon&uid=23028350&db=pubmed&nlmid=101239074"
|
||
title="See full text options at Public Library of Science"
|
||
|
||
data-ga-category="full_text"
|
||
data-ga-action="Public Library of Science"
|
||
data-ga-label="23028350"
|
||
><img src="https://cdn.ncbi.nlm.nih.gov/corehtml/query/egifs/https:--journals.plos.org-resource-img-external-pgen_120x30.png" alt="Public Library of Science full text link"><span class="text">
|
||
|
||
Public Library of Science
|
||
|
||
</span></a><a class="link-item
|
||
pmc
|
||
"
|
||
href="https://pmc.ncbi.nlm.nih.gov/articles/pmid/23028350/"
|
||
target="_blank"
|
||
rel="noopener"
|
||
|
||
ref="linksrc=fulltextorjournal_fulltext&is_pmc=True&PrId=3494&itool=Abstract-def&log$=linkouticon&uid=23028350&db=pubmed&nlmid=101239074"
|
||
title="Free full text at PubMed Central"
|
||
|
||
data-ga-category="full_text"
|
||
data-ga-action="PMC"
|
||
data-ga-label="23028350"
|
||
><span class="text">
|
||
|
||
Free PMC article
|
||
|
||
</span></a>
|
||
</div>
|
||
|
||
</div>
|
||
|
||
<div class="short-view">
|
||
<a href="#" class="full-text-links-button full-text-links-dialog-trigger">
|
||
Full text links
|
||
</a>
|
||
</div>
|
||
</div>
|
||
|
||
|
||
|
||
|
||
<div class="actions-buttons sidebar"><h3 class="title">Actions</h3><div class="inner-wrap"><button class="citation-button citation-dialog-trigger"
|
||
aria-label="Open dialog with citation text in different styles"
|
||
data-ga-category="save_share"
|
||
data-ga-action="cite"
|
||
data-ga-label="open"
|
||
data-all-citations-url="/23028350/citations/"
|
||
data-citation-style="nlm"
|
||
data-pubmed-format-link="/23028350/export/"><span class="button-label">Cite</span></button><link type="text/css" href="ncbi-overlay-block/src/overlay-block.css"><div class="collections-button-container" data-article-id="23028350" data-article-db="pubmed"><button class="collections-button collections-dialog-trigger"
|
||
aria-label="Save article in MyNCBI collections."
|
||
data-ga-category="collections_button"
|
||
data-ga-action="click"
|
||
data-ga-label="collections_button"
|
||
data-collections-open-dialog-enabled="false"
|
||
data-collections-open-dialog-url="https://account.ncbi.nlm.nih.gov/?back_url=https%3A%2F%2Fpubmed.ncbi.nlm.nih.gov%2F23028350%2F%23open-collections-dialog"
|
||
data-in-collections="false"><span class="button-label">Collections</span></button><div class="overlay" role="dialog"><div id="collections-action-dialog"
|
||
class="dialog collections-dialog"
|
||
aria-hidden="true"><div class="title">Add to Collections</div><div class="collections-action-panel action-panel"><form id="collections-action-dialog-form"
|
||
class="collections-action-panel-form action-panel-content action-form action-panel-smaller-selectors"
|
||
data-existing-collections-url="/list-existing-collections/"
|
||
data-add-to-existing-collection-url="/add-to-existing-collection/"
|
||
data-create-and-add-to-new-collection-url="/create-and-add-to-new-collection/"
|
||
data-myncbi-max-collection-name-length="100"
|
||
data-collections-root-url="https://www.ncbi.nlm.nih.gov/myncbi/collections/"><input type="hidden" name="csrfmiddlewaretoken" value="vzMy7zZkDk3vgebaPSut8k1jcJTdfjRPMmwDKpsGdVQZaFh0sb7vCsxXKNBp8Up9"><div class="choice-group" role="radiogroup"><ul class="radio-group-items"><li><input type="radio"
|
||
id="collections-action-dialog-new"
|
||
class="collections-new"
|
||
name="collections"
|
||
value="new"
|
||
data-ga-category="collections_button"
|
||
data-ga-action="click"
|
||
data-ga-label="collections_radio_new"><label for="collections-action-dialog-new">Create a new collection</label></li><li><input type="radio"
|
||
id="collections-action-dialog-existing"
|
||
class="collections-existing"
|
||
name="collections"
|
||
value="existing"
|
||
checked="true"
|
||
data-ga-category="collections_button"
|
||
data-ga-action="click"
|
||
data-ga-label="collections_radio_existing"><label for="collections-action-dialog-existing">Add to an existing collection</label></li></ul></div><div class="controls-wrapper"><div class="action-panel-control-wrap new-collections-controls"><label for="collections-action-dialog-add-to-new" class="action-panel-label required-field-asterisk">
|
||
Name your collection:
|
||
</label><input
|
||
type="text"
|
||
name="add-to-new-collection"
|
||
id="collections-action-dialog-add-to-new"
|
||
class="collections-action-add-to-new"
|
||
pattern="[^"&=<>\/]*" title="The following characters are not allowed in the Name field: "&=<>/"
|
||
maxlength="100"
|
||
data-ga-category="collections_button"
|
||
data-ga-action="create_collection"
|
||
data-ga-label="non_favorties_collection"><div class="collections-new-name-too-long usa-input-error-message selection-validation-message">
|
||
Name must be less than 100 characters
|
||
</div></div><div class="action-panel-control-wrap existing-collections-controls"><label for="collections-action-dialog-add-to-existing" class="action-panel-label">
|
||
Choose a collection:
|
||
</label><select id="collections-action-dialog-add-to-existing"
|
||
class="action-panel-selector collections-action-add-to-existing"
|
||
data-ga-category="collections_button"
|
||
data-ga-action="select_collection"
|
||
data-ga-label="($('.collections-action-add-to-existing').val() === 'Favorites') ? 'Favorites' : 'non_favorites_collection'"></select><div class="collections-retry-load-on-error usa-input-error-message selection-validation-message">
|
||
Unable to load your collection due to an error<br><a href="#">Please try again</a></div></div></div><div class="action-panel-actions"><button class="action-panel-submit"
|
||
type="submit"
|
||
data-loading-label="Adding..."
|
||
data-pinger-ignore
|
||
data-ga-category="collections_button"
|
||
data-ga-action="click"
|
||
data-ga-label="add">
|
||
Add
|
||
</button><button class="action-panel-cancel"
|
||
aria-label="Close 'Add to Collections' panel"
|
||
ref="linksrc=close_collections_panel"
|
||
aria-controls="collections-action-panel"
|
||
aria-expanded="false"
|
||
data-ga-category="collections_button"
|
||
data-ga-action="click"
|
||
data-ga-label="cancel">
|
||
Cancel
|
||
</button></div></form></div></div></div></div><button class="more-actions-button more-actions-dialog-trigger"
|
||
title="Open dialog with more actions to take"
|
||
ref="linksrc=more_actions_btn"></button><div class="display-options" data-format-key="format"
|
||
data-default-format-value="abstract"
|
||
data-current-format-value="abstract" role="region"><button
|
||
class="trigger usa-button"
|
||
title="Change format"
|
||
data-ga-category="display_options"
|
||
data-ga-action="display_options_dialog"><span class="button-label">Display options</span></button><div class="dropdown" hidden><div class="title">
|
||
Display options
|
||
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</a></li></ul></div></div></div><span class="period">. </span><span class="cit">2012 Sep;8(9):e1002938.</span>
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An essential role of variant histone H3.3 for ectomesenchyme potential of the cranial neural crest
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An essential role of variant histone H3.3 for ectomesenchyme potential of the cranial neural crest
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<span class="authors-list-item "><span class="full-name">Samuel G Cox</span><span class="citation-separator"> et al.</span></span>
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The neural crest (NC) is a vertebrate-specific cell population that exhibits remarkable multipotency. Although derived from the neural plate border (NPB) ectoderm, cranial NC (CNC) cells contribute not only to the peripheral nervous system but also to the ectomesenchymal precursors of the head skeleton. To date, the developmental basis for such broad potential has remained elusive. Here, we show that the replacement histone H3.3 is essential during early CNC development for these cells to generate ectomesenchyme and head pigment precursors. In a forward genetic screen in zebrafish, we identified a dominant D123N mutation in h3f3a, one of five zebrafish variant histone H3.3 genes, that eliminates the CNC-derived head skeleton and a subset of pigment cells yet leaves other CNC derivatives and trunk NC intact. Analyses of nucleosome assembly indicate that mutant D123N H3.3 interferes with H3.3 nucleosomal incorporation by forming aberrant H3 homodimers. Consistent with CNC defects arising from insufficient H3.3 incorporation into chromatin, supplying exogenous wild-type H3.3 rescues head skeletal development in mutants. Surprisingly, embryo-wide expression of dominant mutant H3.3 had little effect on embryonic development outside CNC, indicating an unexpectedly specific sensitivity of CNC to defects in H3.3 incorporation. Whereas previous studies had implicated H3.3 in large-scale histone replacement events that generate totipotency during germ line development, our work has revealed an additional role of H3.3 in the broad potential of the ectoderm-derived CNC, including the ability to make the mesoderm-like ectomesenchymal precursors of the head skeleton.
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<p> Figure 1. A dominant H3.3 mutation results in losses of CNC–derived head skeleton and pigment… </p>
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Figure 1. A dominant H3.3 mutation results in losses of CNC–derived head skeleton and pigment cells.
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<div class="figure-caption-contents">a, b, <i>fli1a</i>:GFP-labeled arch ectomesenchyme (arrowheads) is greatly reduced, yet <i>fli1a</i>:GFP-positive endothelial cells (top) are unaffected, in both homozygous and heterozygous <i>h3f3a<sup>db1092</sup></i> mutants at 34 hpf (7/7 mutant; 0/10 wild-type). c, d, Homozygous <i>h3f3a<sup>db1092/db1092</sup></i> embryos specifically lack the CNC-derived head skeleton at 5 dpf (36/71 complete loss; 35/71 partial loss). Diagrams show the CNC-derived cartilage (blue) and bone and teeth (red), mesoderm-derived cartilage (green), pectoral fin cartilage (black), and eyes (yellow). e, f, <i>h3f3a<sup>db1092/+</sup></i> heterozygous larvae exhibit a wide range of craniofacial defects. In some cases, no defects are observed in the facial skeleton and heterozygotes are adult viable (not shown). In mild cases (e), dorsal cartilage and bone of the first and second arches are preferentially reduced, including the dorsal hyosymplectic cartilage and opercular bone of the second arch (arrow). In more severe cases (f), the cartilage and bone of the first arch and dorsal second arch are greatly reduced, with the anterior neurocranium and the posterior ceratobranchial cartilages being less affected. The frequency of skeletal phenotypes in <i>h3f3a<sup>db1092/+</sup></i> heterozygous larvae is highly variable between clutches. g, h, At 27 hpf, <i>dct</i>-positive melanophore precursors are selectively missing anterior to the ear (arrowheads) in both homozygous and heterozygous <i>h3f3a<sup>db1092</sup></i> embryos (5/5 mutant; 0/5 wild-type). i, j, At 27 hpf, cranial <i>xdh</i>-positive xanthophore precursors are mildly reduced in both homozygous and heterozygous <i>h3f3a<sup>db1092</sup></i> embryos (6/8 mutant; 0/4 wild-type). k, l, Wild-type and both homozygous and heterozygous <i>h3f3a<sup>db1092</sup></i> embryos have comparable numbers of <i>foxd3</i>-positive glial cells at 24 hpf (4 mutant; 5 wild-type). m-r, D123N <i>h3f3a</i> mRNA-injected but not wild-type <i>h3f3a</i> mRNA-injected embryos lack <i>fli1a</i>:GFP-positive ectomesenchyme (9/19 D123N; 0/12 wild-type), CNC-derived head skeleton (24/45 D123N; 0/21 wild-type) and cranial <i>dct</i>-positive melanophore precursors (anterior to the ear: arrowheads) (7/14 D123N; 0/11 wild-type). <i>fli1a</i>:GFP-positive blood vessels are unaffected (arrows). s, Except for the loss of the majority of the skull (no facial structures below the level of the eye: arrowheads) and mild heart edema (arrows), the overall morphologies of wild-type, homozygous and heterozygous <i>h3f3a<sup>db1092</sup></i>, wild-type <i>h3f3a</i>-injected, and D123N <i>h3f3a</i>-injected larvae are indistinguishable at 5 dpf. Melanophores (black) and xanthophores (yellow) are also largely normal. Except for panels e and f, homozygous <i>h3f3a<sup>db1092</sup></i> examples are shown. Scale bars: a, b, m & n, 50 µm; c–l, o–s, 250 µm.</div>
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<p> Figure 2. Identification of the <i> h3f3a <sup> db1092… </sup> </i> </p>
|
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<p> Figure 2. Identification of the <i> h3f3a <sup> db1092 </sup> </i> lesion. </p>
|
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<div class="figure-caption-contents"><p> a, The <i> db1092 </i> allele was crossed to the… </p></div>
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<strong class="figure-label">
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Figure 2. Identification of the <i>h3f3a<sup>db1092</sup></i> lesion.
|
||
</strong>
|
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<div class="figure-caption-contents">a, The <i>db1092</i> allele was crossed to the highly polymorphic WIK strain for linkage analysis. As the <i>db1092</i> mutation is semi-dominant, we enriched for putative heterozygotes by selecting for partial head skeletal loss, and events were scored as recombination only if both chromosomes displayed the wild-type WIK polymorphism. Using a set of microsatellite ‘Z’ markers spanning the zebrafish genome, we placed <i>db1092</i> on linkage group 3 near Z3725 and Z20058, and subsequent linkage analysis placed it between Z63643 and Z66457. Recombinants per 1065 meioses are listed above each marker. Sequencing of 3′ UTRs identified single nucleotide polymorphisms (SNPs) that created or destroyed restriction sites between the mutant and WIK chromosomes. These SNPs (identified by their position in millions of base pairs) and Z48485 were then used to map <i>db1092</i> to a 464 kb interval. b, Electrophoretograms show a G to A transition in the <i>h3f3a</i> gene of <i>db1092</i> homozygotes. c, Schematic of the H3.3 variant histone protein encoded by <i>h3f3a</i>. The <i>db1092</i> mutation results in a D123N substitution near the C-terminus of the core domain.</div>
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<p> Figure 3. <i> h3f3a </i> is ubiquitously expressed throughout… </p>
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<p> Figure 3. <i> h3f3a </i> is ubiquitously expressed throughout embryogenesis. </p>
|
||
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<div class="figure-caption-contents"><p> a–i, Lack of <i> h3f3a </i> expression at the… </p></div>
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Figure 3. <i>h3f3a</i> is ubiquitously expressed throughout embryogenesis.
|
||
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|
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<div class="figure-caption-contents">a–i, Lack of <i>h3f3a</i> expression at the one-cell stage shows that <i>h3f3a</i> mRNA is not maternally provided. From 4–14.5 hpf, <i>h3f3a</i> is expressed ubiquitously throughout the embryo. By 16.5 hpf and 27 hpf, <i>h3f3a</i> expression is still widespread, with higher levels apparent in the anterior part of the embryo, including the CNC-derived ectomesenchyme of the pharyngeal arches (arrows) at 27 hpf. Scale bars = 250 µm.</div>
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<p> Figure 4. The dominant D123N mutation prevents… </p>
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||
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<p> Figure 4. The dominant D123N mutation prevents chromatin incorporation and promotes the formation of aberrant… </p>
|
||
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|
||
Figure 4. The dominant D123N mutation prevents chromatin incorporation and promotes the formation of aberrant H3 homodimers.
|
||
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|
||
|
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|
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|
||
<div class="figure-caption-contents">a, Western blots show α-FLAG immunostaining of nuclear extracts or purified mononucleosome fractions from HEK cells transfected with vector alone or FLAG-tagged H3.3 (f:H3.3) vectors. Wild-type and D123N f:H3.3 proteins are expressed at equal levels in total extract, but wild-type f:H3.3 is present at much higher levels in the nucleosome fraction (consistent over three replicate experiments). α-FLAG immunoprecipitation from purified nucleosomes shows that wild-type but not D123N f:H3.3 is incorporated into nucleosomes containing H2A, H2B, H3, and H4 (consistent over three replicate experiments). b, Confocal images from H2A.F/Z:GFP embryos expressing wild-type and D123N versions of mCherry(m)H3.3 and mCherry(m)H3.2 fusion proteins. Merged images show that whereas all H3 proteins are nuclear localized in surrounding non-mitotic cells, wild-type mH3.3 and mH3.2, but not D123N mH3.3 and mH3.2, co-localize with H2A.F/Z:GFP in the chromosomes of metaphase/anaphase cells (arrowheads) after nuclear envelope breakdown (wild-type mH3.3, 11/11 cells in 2 embryos; D123N mH3.3, 0/25 cells in 3 embryos; wild-type mH3.2, 21/21 cells in 3 embryos; D123N mH3.2, 0/16 cells in 2 embryos). c, α-FLAG, α-H3 and α-H4 western blots for samples immunoprecipitated by α-FLAG from nuclear extracts of f:H3.3-transfected HEK cells. Recombinant octamer is used as a reference. Whereas both endogenous H3 and H4 co-immunoprecipitate with the wild-type f:H3.3 protein (*), H3 but not H4 co-immunoprecipitates with D123N f:H3.3 (asterisk marks the larger recombinant f:H3.3 protein). Results were consistent over three replicate experiments. d, mRNA injection of D123N mH3.3 (8/17), but not wild-type mH3.3 (0/26), wild-type mH3.2 (0/19), or D123N mH3.2 (0/18), results in loss of the CNC-derived head skeleton at 4 dpf. Scale bars: b, 10 µm; d, 250 µm.</div>
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<p> Figure 5. Injection of wild-type H3.3 RNA… </p>
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<p> Figure 5. Injection of wild-type H3.3 RNA and reduction of mutant H3.3 levels both rescue… </p>
|
||
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|
||
Figure 5. Injection of wild-type H3.3 RNA and reduction of mutant H3.3 levels both rescue craniofacial skeletal development in <i>h3f3a<sup>db1092</sup></i> mutants.
|
||
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|
||
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|
||
<div class="figure-caption-contents">a, b, Results from scoring the severity of craniofacial losses in 5 dpf larval head skeletons from <i>h3f3a<sup>db1092</sup></i> siblings injected with 450 ng/µl mRNA encoding wild-type H3.3 or a control kikGR fluorescent protein. The scoring system ranges from Grade 0 (wild-type phenotype) to Grade 5 (complete loss of CNC derivatives); see Materials and Methods for more detail. H3.3-mRNA-injected <i>h3f3a<sup>db1092</sup></i> homozygotes (a) and heterozygotes (b) exhibited a decrease in the severity of <i>h3f3a<sup>db1092</sup></i> craniofacial phenotypes over kikGR-RNA-injected controls (significant by Fisher's exact test: homozygotes, p = 7.7E-05; heterozygotes, p = 1.0E-04). c, An antisense morpholino oligonucleotide was designed to inhibit splicing at the exon 3/intron 3–4 boundary (green arrowhead) of the <i>h3f3a</i> transcript. Morpholinos were injected into one-cell-stage <i>h3f3a<sup>db1092</sup></i> embryos at 400 µM. d, Morpholino efficacy was demonstrated by PCR amplification between exons flanking the targeted splice junction from 10 hpf cDNA from 20 pooled embryos (position of primers shown as red arrows in c). Compared to the sample from uninjected (un) embryos, the morpholino-treated sample (MO) exhibited a partial decrease in PCR product representing spliced transcript (295 bp) and a concomitant increase in un-spliced PCR product (390 bp). e, f, Compared to uninjected siblings, both morpholino-injected <i>h3f3a<sup>db1092</sup></i> homozygotes (e) and heterozygotes (f) exhibited a decrease in the severity of <i>h3f3a<sup>db1092</sup></i> craniofacial phenotypes (significant by Fisher's exact test: homozygotes, p = 2.0E-04; heterozygotes, p = 6.3E-06). (g) Craniofacial development in wild-type embryos was unaffected by morpholino injection.</div>
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<p> Figure 6. H3.3 functions at the NPB–CNC… </p>
|
||
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<p> Figure 6. H3.3 functions at the NPB–CNC transition. </p>
|
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<div class="figure-caption-contents"><p> a–f, Expression of <i> msxb </i> , <i> pax3a </i> ,… </p></div>
|
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Figure 6. H3.3 functions at the NPB–CNC transition.
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<div class="figure-caption-contents">a–f, Expression of <i>msxb</i>, <i>pax3a</i>, <i>zic2a</i>, and <i>tfap2a</i> at 10 hpf and <i>msxb</i> and <i>pax3a</i> at 11 hpf is indistinguishable between wild types and both homozygous and heterozygous <i>h3f3a<sup>db1092</sup></i> mutants (mut) (n≥10 for each). g–k, At 11 hpf, both homozygous and heterozygous <i>h3f3a<sup>db1092</sup></i> mutants have severe reductions in the expression of <i>snai2</i> (4/4 mut; 0/4 wt), <i>sox10</i> (10/12 mut; 0/6 wild-type), <i>foxd3</i> (9/9 mut; 0/5 wt), <i>tfap2a</i> (7/7 mut; 0/4 wt), and <i>sox9b</i> (8/8 mut; 0/3 wt). l, <i>sox10</i> expression is also lost in embryos injected with D123N (10/12) but not wild-type (0/16) <i>h3f3a</i> mRNA. m, In both homozygous and heterozygous <i>h3f3a<sup>db1092</sup></i> embryos, <i>sox10</i> expression partially recovers by 16.5 hpf yet is specifically reduced in presumptive CNC ectomesenchyme domains (arrows) (6/6 mut; 0/4 wt). An increase in <i>sox10</i>-positive cells is evident in the mutant dorsal neural tube (insert) between the <i>sox10</i>-positive otic placodes (arrowheads) which are unaffected in mutants. n, At 16.5 hpf, <i>dlx2a</i> expression in three streams of migrating ectomesenchyme is reduced in both homozygous and heterozygous <i>h3f3a<sup>db1092</sup></i> mutants (6/7 mut; 0/5 wt). o, The 16.5 hpf ectomesenchyme expression (arrows) of <i>twist1a</i> is reduced in both homozygous and heterozygous <i>h3f3a<sup>db1092</sup></i> mutants yet paraxial mesoderm expression is unaffected (white arrowheads) (8/8 mut; 0/5 wt). In all panels, homozygous <i>h3f3a<sup>db1092</sup></i> examples are shown. All images are dorsal views with anterior to the left. Scale bars: 250 µm.</div>
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<p> Figure 7. Trunk NC is largely unaffected… </p>
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<p> Figure 7. Trunk NC is largely unaffected in <i> h3f3a <sup> db1092 </sup> </i> mutants. </p>
|
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<div class="figure-caption-contents"><p> a–c, <i> crestin </i> expression at 11.7… </p></div>
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Figure 7. Trunk NC is largely unaffected in <i>h3f3a<sup>db1092</sup></i> mutants.
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<div class="figure-caption-contents">a–c, <i>crestin</i> expression at 11.7 hpf shows similar amounts of trunk NC in wild-type and both homozygous and heterozygous <i>h3f3a<sup>db1092</sup></i> embryos (n = 5 for each genotype). d–i, Trunk views of <i>sox9b</i> expression at 11.7 hpf show that trunk NC specification is largely normal in both homozygous and heterozygous <i>h3f3a<sup>db1092</sup></i> mutants (n = 10 for each genotype). Cranial views of the same embryos show reduced amounts of <i>sox9b</i>-expressing CNC. Arrows show the <i>sox9b</i>-positive otic placodes that are unaffected in mutants. Scale bars = 250 µm.</div>
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<p> Figure 8. H3.3 function is required tissue-… </p>
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<p> Figure 8. H3.3 function is required tissue- and cell-autonomously for CNC development. </p>
|
||
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<div class="figure-caption-contents"><p> a, Wild-type cells… </p></div>
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Figure 8. H3.3 function is required tissue- and cell-autonomously for CNC development.
|
||
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|
||
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|
||
<div class="figure-caption-contents">a, Wild-type cells were transplanted unilaterally into the CNC precursor domain of <i>h3f3a<sup>db1092/db1092</sup></i> homozygous mutants at 6 hpf. b, Compared to the non-recipient control side (bottom), expression of the early CNC marker <i>snai2</i> is restored in the recipient side at 11hpf (top) (n = 17/29 with rescue). c–e, <i>fli1a</i>:GFP (green) marks CNC ectomesenchyme of the pharyngeal arches at 30 hpf and facial skeletal elements at 5 dpf. The red fluorescent dye, Alexa568, marks transplanted wild-type cells, whereas both donor and host cells harbor the <i>fli1a</i>:GFP transgene. When transplanted into an <i>h3f3a<sup>db1092/db1092</sup></i> homozygous host, wild-type Alexa568+ CNC precursors contribute to pharyngeal arch ectomesenchyme and rescue arch size (c) and form wild-type cartilage and bone (e). In contrast, the non-recipient control side (d) has reduced pharyngeal arch ectomesenchyme. Whereas wild-type donor cells appear yellow due to red Alexa568 and green <i>fli1a</i>:GFP, mutant host cells have only <i>fli1a</i>:GFP and hence appear green. Rescue of arch size was observed in 8/11 cases. f, Alcian staining at 5 dpf shows that cartilage is restored to half the face in an <i>h3f3a<sup>db1092/db1092</sup></i> larvae that received an unilateral wild-type CNC precursor transplant. Compare the recipient side (left) to the control side that forms little facial cartilage (right). Skeletal rescue was observed in 21/30 cases. g, Individual cells of 32-cell stage <i>sox10</i>:GFP embryos were injected with mRNA encoding mCherry-tagged versions of wild-type or D123N H3.3 to generate mosaic mCherry-H3.3 expression at later stages. h, Soon after the appearance of GFP-labeled CNC at approximately 11 hpf, mosaic embryos were assessed for incorporation of mCherry-H3.3-expressing cells (red) into the <i>sox10</i>:GFP-positive CNC domain (green). i/i′/i″ and j/j′/j″, Confocal images from <i>sox10</i>:GFP embryos with mosaic expression of wild-type (i/i′/i″) and D123N (j/j′/j″) versions of mCherry-H3.3. Cells doubly-positive for wild-type mCherry-H3.3 (red) and GFP (green) (arrowheads) were observed within the CNC domain (7/14 cells over 4 embryos), whereas mutant D123N mCherry-H3.3 cells within the CNC domain failed to up-regulate <i>sox10</i>:GFP (arrowheads) (0/18 cells over 4 embryos). Only cells with strong mCherry-H3.3 were used in the analysis. hs: hyosymplectic cartilage, pq: palatoquadrate cartilage, ch: ceratohyal cartilage, op: opercular bone. Scale bars: b, f, 250 µm; c–e, 50 µm; i and j,10 µm.</div>
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<p> Figure 9. Cell death in <i> h3f3a <sup> db1092… </sup> </i> </p>
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<p> Figure 9. Cell death in <i> h3f3a <sup> db1092 </sup> </i> embryos. </p>
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Figure 9. Cell death in <i>h3f3a<sup>db1092</sup></i> embryos.
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<div class="figure-caption-contents">a–c, Lysotracker Red staining marks similar amounts of dying cells in wild-type (n = 2), <i>h3f3a<sup>db1092/+</sup></i> heterozygous (n = 5), and <i>h3f3a<sup>db1092/db1092</sup></i> homozygous (n = 3) embryos at 12.5 hpf. The bright staining in the bottom of each panel is the yolk. d–f, At 16 hpf, increased Lysotracker Red staining (arrows) was evident in the dorsal neural tube of <i>h3f3a<sup>db1092/+</sup></i> heterozygotes (2/2) and <i>h3f3a<sup>db1092/db1092</sup></i> homozygotes (3/3) but not wild types (0/4). These dying cells were located in a similar position to where CNC forms in wild-type embryos. Scale bar = 50 µm.</div>
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<p> Figure 10. Model for the role of H3.3-dependent histone replacement during CNC development. </p>
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Figure 10. Model for the role of H3.3-dependent histone replacement during CNC development.
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<div class="figure-caption-contents">At the early embryonic blastula stage, cells have a broad potential with cis-regulatory elements for developmental genes existing in a “poised” chromatin state. After gastrulation occurs to form the three major germ layers (ectoderm, mesoderm, and endoderm), genes associated with a particular germ layer are activated or maintained in a poised state, whereas genes for other layers are strongly repressed at the chromatin level. The cranial neural crest (CNC) is unusual in that it is derived from ectoderm yet can give rise to mesoderm-like derivatives such as skeleton. H3.3-dependent histone replacement could thus be required to remodel the enhancers of mesodermal genes needed for ectomesenchymal fates, with the distinctive role of H3.3 in CNC correlating with the need to derepress mesodermal enhancers that have been previously silenced in the ectoderm germ layer (1). Alternatively H3.3 incorporation could act to maintain mesoderm-like potential in the CNC ectoderm from an earlier time in development (2). It also remains unresolved the extent to which ectomesenchyme derivatives (e.g. head skeleton) and non-ectomesenchyme derivatives (e.g. pigment, glia, and neurons) derive from a common multipotent precursor. Hence, the cranial pigment and ectomesenchyme defects of <i>h3f3a<sup>db1092</sup></i> mutants could arise from altered histone replacement in a common multipotent precursor, or alternatively from independent defects in different subsets of heterogeneous CNC with more limited potential.</div>
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Baroffio A, Dupin E, Le Douarin NM (1991) Common precursors for neural and mesectodermal derivatives in the cephalic neural crest. Development 112: 301–305.
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Bronner-Fraser M, Fraser SE (1988) Cell lineage analysis reveals multipotency of some avian neural crest cells. Nature 335: 161–164.
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Platt JB (1893) Ectodermic origin of the cartilages of the head. Anat Anz 8: 506–509.
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Schilling TF, Kimmel CB (1994) Segment and cell type lineage restrictions during pharyngeal arch development in the zebrafish embryo. Development 120: 483–494.
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Research Support, Non-U.S. Gov't
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|
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</a></li></ul></div></div></div></li><li><div class="keyword-actions dropdown-block"><button class="keyword-actions-trigger trigger keyword-link"
|
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aria-label="Toggle dropdown menu for keyword Embryonic Development / genetics"
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aria-controls="keyword-actions-mesh-terms-7"
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aria-expanded="false"
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Embryonic Development / genetics
|
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</button><div id="keyword-actions-mesh-terms-7"
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class="keyword-actions-dropdown dropdown dropdown-container"
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hidden
|
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aria-label="Dropdown menu for keyword Embryonic Development / genetics"><div class="title">Actions</div><div class="content"><ul class="keyword-actions-links"><li><a class="search-in-pubmed-link dropdown-block-link"
|
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href="/?term=%22Embryonic+Development%2Fgenetics%22%5BMeSH%5D&sort=date&sort_order=desc"
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data-href="/?term=%22Embryonic+Development%2Fgenetics%22%5BMeSH%5D&sort=date&sort_order=desc"
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ref="linksrc=search_in_pubmed_keyword_link&mesh="Embryonic Development/genetics"[MeSH]"
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</a></li><li><a class="add-to-search-link dropdown-block-link"
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</a></li></ul></div></div></div></li><li><div class="keyword-actions dropdown-block"><button class="keyword-actions-trigger trigger keyword-link"
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aria-label="Toggle dropdown menu for keyword Gene Expression Regulation, Developmental"
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Gene Expression Regulation, Developmental
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</button><div id="keyword-actions-mesh-terms-8"
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aria-label="Dropdown menu for keyword Gene Expression Regulation, Developmental"><div class="title">Actions</div><div class="content"><ul class="keyword-actions-links"><li><a class="search-in-pubmed-link dropdown-block-link"
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href="/?term=%22Gene+Expression+Regulation%2C+Developmental%22%5BMeSH%5D&sort=date&sort_order=desc"
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data-href="/?term=%22Gene+Expression+Regulation%2C+Developmental%22%5BMeSH%5D&sort=date&sort_order=desc"
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</a></li></ul></div></div></div></li><li><div class="keyword-actions dropdown-block"><button class="keyword-actions-trigger trigger keyword-link"
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aria-label="Dropdown menu for keyword HEK293 Cells"><div class="title">Actions</div><div class="content"><ul class="keyword-actions-links"><li><a class="search-in-pubmed-link dropdown-block-link"
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</a></li></ul></div></div></div></li><li><div class="keyword-actions dropdown-block"><button class="keyword-actions-trigger trigger keyword-link major"
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aria-label="Toggle dropdown menu for keyword Histones / genetics*"
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Histones / genetics*
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aria-label="Dropdown menu for keyword Histones / genetics*"><div class="title">Actions</div><div class="content"><ul class="keyword-actions-links"><li><a class="search-in-pubmed-link dropdown-block-link"
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href="/?term=%22Histones%2Fgenetics%22%5BMAJR%5D&sort=date&sort_order=desc"
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</a></li></ul></div></div></div></li><li><div class="keyword-actions dropdown-block"><button class="keyword-actions-trigger trigger keyword-link"
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aria-label="Toggle dropdown menu for keyword Histones / metabolism"
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Histones / metabolism
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aria-label="Dropdown menu for keyword Histones / metabolism"><div class="title">Actions</div><div class="content"><ul class="keyword-actions-links"><li><a class="search-in-pubmed-link dropdown-block-link"
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href="/?term=%22Histones%2Fmetabolism%22%5BMeSH%5D&sort=date&sort_order=desc"
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data-href="/?term=%22Histones%2Fmetabolism%22%5BMeSH%5D&sort=date&sort_order=desc"
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</a></li></ul></div></div></div></li><li><div class="keyword-actions dropdown-block"><button class="keyword-actions-trigger trigger keyword-link"
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aria-label="Toggle dropdown menu for keyword Humans"
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Humans
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</button><div id="keyword-actions-mesh-terms-12"
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class="keyword-actions-dropdown dropdown dropdown-container"
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aria-label="Dropdown menu for keyword Humans"><div class="title">Actions</div><div class="content"><ul class="keyword-actions-links"><li><a class="search-in-pubmed-link dropdown-block-link"
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href="/?term=%22Humans%22%5BMeSH%5D&sort=date&sort_order=desc"
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data-href="/?term=%22Humans%22%5BMeSH%5D&sort=date&sort_order=desc"
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</a></li></ul></div></div></div></li><li><div class="keyword-actions dropdown-block"><button class="keyword-actions-trigger trigger keyword-link"
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aria-label="Toggle dropdown menu for keyword Mesoderm / growth & development"
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aria-expanded="false"
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Mesoderm / growth & development
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</button><div id="keyword-actions-mesh-terms-13"
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class="keyword-actions-dropdown dropdown dropdown-container"
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hidden
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aria-label="Dropdown menu for keyword Mesoderm / growth & development"><div class="title">Actions</div><div class="content"><ul class="keyword-actions-links"><li><a class="search-in-pubmed-link dropdown-block-link"
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href="/?term=%22Mesoderm%2Fgrowth+and+development%22%5BMeSH%5D&sort=date&sort_order=desc"
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data-href="/?term=%22Mesoderm%2Fgrowth+and+development%22%5BMeSH%5D&sort=date&sort_order=desc"
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ref="linksrc=search_in_pubmed_keyword_link&mesh="Mesoderm/growth and development"[MeSH]"
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Search in MeSH
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Add to Search
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</a></li></ul></div></div></div></li><li><div class="keyword-actions dropdown-block"><button class="keyword-actions-trigger trigger keyword-link"
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aria-label="Toggle dropdown menu for keyword Mutation"
|
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tabindex="0"
|
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aria-controls="keyword-actions-mesh-terms-14"
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aria-expanded="false"
|
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data-pinger-ignore>
|
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Mutation
|
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</button><div id="keyword-actions-mesh-terms-14"
|
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class="keyword-actions-dropdown dropdown dropdown-container"
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hidden
|
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aria-label="Dropdown menu for keyword Mutation"><div class="title">Actions</div><div class="content"><ul class="keyword-actions-links"><li><a class="search-in-pubmed-link dropdown-block-link"
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href="/?term=%22Mutation%22%5BMeSH%5D&sort=date&sort_order=desc"
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data-href="/?term=%22Mutation%22%5BMeSH%5D&sort=date&sort_order=desc"
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ref="linksrc=search_in_pubmed_keyword_link&mesh="Mutation"[MeSH]"
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href="https://www.ncbi.nlm.nih.gov/mesh?term=Mutation"
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Search in MeSH
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</a></li><li><a class="add-to-search-link dropdown-block-link"
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data-search-term=""Mutation"[MeSH]"
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data-ga-label=""Mutation"[MeSH]"
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Add to Search
|
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</a></li></ul></div></div></div></li><li><div class="keyword-actions dropdown-block"><button class="keyword-actions-trigger trigger keyword-link"
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aria-label="Toggle dropdown menu for keyword Neural Crest / cytology"
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tabindex="0"
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aria-expanded="false"
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Neural Crest / cytology
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</button><div id="keyword-actions-mesh-terms-15"
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hidden
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aria-label="Dropdown menu for keyword Neural Crest / cytology"><div class="title">Actions</div><div class="content"><ul class="keyword-actions-links"><li><a class="search-in-pubmed-link dropdown-block-link"
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data-href="/?term=%22Neural+Crest%2Fcytology%22%5BMeSH%5D&sort=date&sort_order=desc"
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Add to Search
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</a></li></ul></div></div></div></li><li><div class="keyword-actions dropdown-block"><button class="keyword-actions-trigger trigger keyword-link major"
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aria-label="Toggle dropdown menu for keyword Neural Crest / growth & development*"
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Neural Crest / growth & development*
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hidden
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aria-label="Dropdown menu for keyword Neural Crest / growth & development*"><div class="title">Actions</div><div class="content"><ul class="keyword-actions-links"><li><a class="search-in-pubmed-link dropdown-block-link"
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href="/?term=%22Neural+Crest%2Fgrowth+and+development%22%5BMAJR%5D&sort=date&sort_order=desc"
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data-href="/?term=%22Neural+Crest%2Fgrowth+and+development%22%5BMAJR%5D&sort=date&sort_order=desc"
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Search in MeSH
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data-search-term=""Neural Crest/growth and development"[MAJR]"
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data-ga-category="search"
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Add to Search
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</a></li></ul></div></div></div></li><li><div class="keyword-actions dropdown-block"><button class="keyword-actions-trigger trigger keyword-link"
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aria-label="Toggle dropdown menu for keyword Neural Crest / metabolism"
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data-pinger-ignore>
|
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Neural Crest / metabolism
|
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</button><div id="keyword-actions-mesh-terms-17"
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class="keyword-actions-dropdown dropdown dropdown-container"
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hidden
|
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aria-label="Dropdown menu for keyword Neural Crest / metabolism"><div class="title">Actions</div><div class="content"><ul class="keyword-actions-links"><li><a class="search-in-pubmed-link dropdown-block-link"
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href="/?term=%22Neural+Crest%2Fmetabolism%22%5BMeSH%5D&sort=date&sort_order=desc"
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data-href="/?term=%22Neural+Crest%2Fmetabolism%22%5BMeSH%5D&sort=date&sort_order=desc"
|
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ref="linksrc=search_in_pubmed_keyword_link&mesh="Neural Crest/metabolism"[MeSH]"
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data-ga-category="search"
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data-ga-action="mesh_term"
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data-ga-label="Neural Crest / metabolism">
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Search in PubMed
|
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</a></li><li><a class="search-in-mesh-link dropdown-block-link"
|
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href="https://www.ncbi.nlm.nih.gov/mesh?term=Neural+Crest"
|
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data-ga-category="search_mesh"
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data-ga-action="mesh_term_link"
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data-ga-label="Neural Crest / metabolism">
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Search in MeSH
|
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</a></li><li><a class="add-to-search-link dropdown-block-link"
|
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data-search-term=""Neural Crest/metabolism"[MeSH]"
|
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data-ga-category="search"
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data-ga-action="add_to_search"
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data-ga-label=""Neural Crest/metabolism"[MeSH]"
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Add to Search
|
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</a></li></ul></div></div></div></li><li><div class="keyword-actions dropdown-block"><button class="keyword-actions-trigger trigger keyword-link"
|
||
aria-label="Toggle dropdown menu for keyword Neural Plate / cytology"
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tabindex="0"
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aria-controls="keyword-actions-mesh-terms-18"
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aria-expanded="false"
|
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data-pinger-ignore>
|
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Neural Plate / cytology
|
||
</button><div id="keyword-actions-mesh-terms-18"
|
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class="keyword-actions-dropdown dropdown dropdown-container"
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hidden
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aria-label="Dropdown menu for keyword Neural Plate / cytology"><div class="title">Actions</div><div class="content"><ul class="keyword-actions-links"><li><a class="search-in-pubmed-link dropdown-block-link"
|
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href="/?term=%22Neural+Plate%2Fcytology%22%5BMeSH%5D&sort=date&sort_order=desc"
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data-href="/?term=%22Neural+Plate%2Fcytology%22%5BMeSH%5D&sort=date&sort_order=desc"
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ref="linksrc=search_in_pubmed_keyword_link&mesh="Neural Plate/cytology"[MeSH]"
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Search in PubMed
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href="https://www.ncbi.nlm.nih.gov/mesh?term=Neural+Plate"
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Search in MeSH
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</a></li><li><a class="add-to-search-link dropdown-block-link"
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data-search-term=""Neural Plate/cytology"[MeSH]"
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Add to Search
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</a></li></ul></div></div></div></li><li><div class="keyword-actions dropdown-block"><button class="keyword-actions-trigger trigger keyword-link"
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aria-label="Toggle dropdown menu for keyword Neural Plate / growth & development"
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aria-expanded="false"
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data-pinger-ignore>
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Neural Plate / growth & development
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</button><div id="keyword-actions-mesh-terms-19"
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class="keyword-actions-dropdown dropdown dropdown-container"
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hidden
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aria-label="Dropdown menu for keyword Neural Plate / growth & development"><div class="title">Actions</div><div class="content"><ul class="keyword-actions-links"><li><a class="search-in-pubmed-link dropdown-block-link"
|
||
href="/?term=%22Neural+Plate%2Fgrowth+and+development%22%5BMeSH%5D&sort=date&sort_order=desc"
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data-href="/?term=%22Neural+Plate%2Fgrowth+and+development%22%5BMeSH%5D&sort=date&sort_order=desc"
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ref="linksrc=search_in_pubmed_keyword_link&mesh="Neural Plate/growth and development"[MeSH]"
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Search in PubMed
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</a></li><li><a class="search-in-mesh-link dropdown-block-link"
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Search in MeSH
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</a></li><li><a class="add-to-search-link dropdown-block-link"
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data-search-term=""Neural Plate/growth and development"[MeSH]"
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data-ga-category="search"
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data-ga-label=""Neural Plate/growth and development"[MeSH]"
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Add to Search
|
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</a></li></ul></div></div></div></li><li><div class="keyword-actions dropdown-block"><button class="keyword-actions-trigger trigger keyword-link"
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aria-label="Toggle dropdown menu for keyword Neural Plate / metabolism"
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tabindex="0"
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aria-expanded="false"
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data-pinger-ignore>
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Neural Plate / metabolism
|
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</button><div id="keyword-actions-mesh-terms-20"
|
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class="keyword-actions-dropdown dropdown dropdown-container"
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hidden
|
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aria-label="Dropdown menu for keyword Neural Plate / metabolism"><div class="title">Actions</div><div class="content"><ul class="keyword-actions-links"><li><a class="search-in-pubmed-link dropdown-block-link"
|
||
href="/?term=%22Neural+Plate%2Fmetabolism%22%5BMeSH%5D&sort=date&sort_order=desc"
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data-href="/?term=%22Neural+Plate%2Fmetabolism%22%5BMeSH%5D&sort=date&sort_order=desc"
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ref="linksrc=search_in_pubmed_keyword_link&mesh="Neural Plate/metabolism"[MeSH]"
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Search in PubMed
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</a></li><li><a class="search-in-mesh-link dropdown-block-link"
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href="https://www.ncbi.nlm.nih.gov/mesh?term=Neural+Plate"
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Search in MeSH
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</a></li><li><a class="add-to-search-link dropdown-block-link"
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data-search-term=""Neural Plate/metabolism"[MeSH]"
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data-ga-category="search"
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data-ga-action="add_to_search"
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data-ga-label=""Neural Plate/metabolism"[MeSH]"
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Add to Search
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</a></li></ul></div></div></div></li><li><div class="keyword-actions dropdown-block"><button class="keyword-actions-trigger trigger keyword-link"
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aria-label="Toggle dropdown menu for keyword Nucleosomes / genetics"
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tabindex="0"
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aria-controls="keyword-actions-mesh-terms-21"
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aria-expanded="false"
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data-pinger-ignore>
|
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Nucleosomes / genetics
|
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</button><div id="keyword-actions-mesh-terms-21"
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class="keyword-actions-dropdown dropdown dropdown-container"
|
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hidden
|
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aria-label="Dropdown menu for keyword Nucleosomes / genetics"><div class="title">Actions</div><div class="content"><ul class="keyword-actions-links"><li><a class="search-in-pubmed-link dropdown-block-link"
|
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href="/?term=%22Nucleosomes%2Fgenetics%22%5BMeSH%5D&sort=date&sort_order=desc"
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data-href="/?term=%22Nucleosomes%2Fgenetics%22%5BMeSH%5D&sort=date&sort_order=desc"
|
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ref="linksrc=search_in_pubmed_keyword_link&mesh="Nucleosomes/genetics"[MeSH]"
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Search in PubMed
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</a></li><li><a class="search-in-mesh-link dropdown-block-link"
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href="https://www.ncbi.nlm.nih.gov/mesh?term=Nucleosomes"
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Search in MeSH
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</a></li><li><a class="add-to-search-link dropdown-block-link"
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data-search-term=""Nucleosomes/genetics"[MeSH]"
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data-ga-category="search"
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Add to Search
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</a></li></ul></div></div></div></li><li><div class="keyword-actions dropdown-block"><button class="keyword-actions-trigger trigger keyword-link major"
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aria-label="Toggle dropdown menu for keyword Skull / growth & development*"
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tabindex="0"
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aria-controls="keyword-actions-mesh-terms-major-22"
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aria-expanded="false"
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data-pinger-ignore>
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Skull / growth & development*
|
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</button><div id="keyword-actions-mesh-terms-major-22"
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hidden
|
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aria-label="Dropdown menu for keyword Skull / growth & development*"><div class="title">Actions</div><div class="content"><ul class="keyword-actions-links"><li><a class="search-in-pubmed-link dropdown-block-link"
|
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href="/?term=%22Skull%2Fgrowth+and+development%22%5BMAJR%5D&sort=date&sort_order=desc"
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data-href="/?term=%22Skull%2Fgrowth+and+development%22%5BMAJR%5D&sort=date&sort_order=desc"
|
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ref="linksrc=search_in_pubmed_keyword_link&mesh="Skull/growth and development"[MAJR]"
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Search in PubMed
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href="https://www.ncbi.nlm.nih.gov/mesh?term=Skull"
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Search in MeSH
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</a></li><li><a class="add-to-search-link dropdown-block-link"
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data-search-term=""Skull/growth and development"[MAJR]"
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data-ga-category="search"
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data-ga-action="add_to_search"
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Add to Search
|
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</a></li></ul></div></div></div></li><li><div class="keyword-actions dropdown-block"><button class="keyword-actions-trigger trigger keyword-link"
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aria-label="Toggle dropdown menu for keyword Skull / metabolism"
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tabindex="0"
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aria-controls="keyword-actions-mesh-terms-23"
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aria-expanded="false"
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data-pinger-ignore>
|
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Skull / metabolism
|
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</button><div id="keyword-actions-mesh-terms-23"
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class="keyword-actions-dropdown dropdown dropdown-container"
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hidden
|
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aria-label="Dropdown menu for keyword Skull / metabolism"><div class="title">Actions</div><div class="content"><ul class="keyword-actions-links"><li><a class="search-in-pubmed-link dropdown-block-link"
|
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href="/?term=%22Skull%2Fmetabolism%22%5BMeSH%5D&sort=date&sort_order=desc"
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data-href="/?term=%22Skull%2Fmetabolism%22%5BMeSH%5D&sort=date&sort_order=desc"
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ref="linksrc=search_in_pubmed_keyword_link&mesh="Skull/metabolism"[MeSH]"
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data-ga-label="Skull / metabolism">
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Search in PubMed
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</a></li><li><a class="search-in-mesh-link dropdown-block-link"
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href="https://www.ncbi.nlm.nih.gov/mesh?term=Skull"
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data-ga-category="search_mesh"
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data-ga-action="mesh_term_link"
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Search in MeSH
|
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</a></li><li><a class="add-to-search-link dropdown-block-link"
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role="button"
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data-search-term=""Skull/metabolism"[MeSH]"
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data-ga-category="search"
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data-ga-action="add_to_search"
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data-ga-label=""Skull/metabolism"[MeSH]"
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Add to Search
|
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</a></li></ul></div></div></div></li><li><div class="keyword-actions dropdown-block"><button class="keyword-actions-trigger trigger keyword-link major"
|
||
aria-label="Toggle dropdown menu for keyword Zebrafish* / genetics"
|
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tabindex="0"
|
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aria-controls="keyword-actions-mesh-terms-major-24"
|
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aria-expanded="false"
|
||
data-pinger-ignore>
|
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Zebrafish* / genetics
|
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</button><div id="keyword-actions-mesh-terms-major-24"
|
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class="keyword-actions-dropdown dropdown dropdown-container"
|
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hidden
|
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aria-label="Dropdown menu for keyword Zebrafish* / genetics"><div class="title">Actions</div><div class="content"><ul class="keyword-actions-links"><li><a class="search-in-pubmed-link dropdown-block-link"
|
||
href="/?term=%22Zebrafish%2Fgenetics%22%5BMAJR%5D&sort=date&sort_order=desc"
|
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data-href="/?term=%22Zebrafish%2Fgenetics%22%5BMAJR%5D&sort=date&sort_order=desc"
|
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ref="linksrc=search_in_pubmed_keyword_link&mesh="Zebrafish/genetics"[MAJR]"
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data-ga-category="search"
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data-ga-action="mesh_term"
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data-ga-label="Zebrafish* / genetics">
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Search in PubMed
|
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</a></li><li><a class="search-in-mesh-link dropdown-block-link"
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ref="linksrc=search_in_mesh_link"
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href="https://www.ncbi.nlm.nih.gov/mesh?term=Zebrafish"
|
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data-ga-category="search_mesh"
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data-ga-action="mesh_term_link"
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data-ga-label="Zebrafish* / genetics">
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Search in MeSH
|
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</a></li><li><a class="add-to-search-link dropdown-block-link"
|
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ref="linksrc=add_to_search_link"
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role="button"
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data-search-term=""Zebrafish/genetics"[MAJR]"
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data-ga-category="search"
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data-ga-action="add_to_search"
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data-ga-label=""Zebrafish/genetics"[MAJR]"
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Add to Search
|
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</a></li></ul></div></div></div></li><li><div class="keyword-actions dropdown-block"><button class="keyword-actions-trigger trigger keyword-link major"
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aria-label="Toggle dropdown menu for keyword Zebrafish* / growth & development"
|
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tabindex="0"
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aria-controls="keyword-actions-mesh-terms-major-25"
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aria-expanded="false"
|
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data-pinger-ignore>
|
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Zebrafish* / growth & development
|
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</button><div id="keyword-actions-mesh-terms-major-25"
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class="keyword-actions-dropdown dropdown dropdown-container"
|
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hidden
|
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aria-label="Dropdown menu for keyword Zebrafish* / growth & development"><div class="title">Actions</div><div class="content"><ul class="keyword-actions-links"><li><a class="search-in-pubmed-link dropdown-block-link"
|
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href="/?term=%22Zebrafish%2Fgrowth+and+development%22%5BMAJR%5D&sort=date&sort_order=desc"
|
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data-href="/?term=%22Zebrafish%2Fgrowth+and+development%22%5BMAJR%5D&sort=date&sort_order=desc"
|
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ref="linksrc=search_in_pubmed_keyword_link&mesh="Zebrafish/growth and development"[MAJR]"
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data-ga-category="search"
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data-ga-action="mesh_term"
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data-ga-label="Zebrafish* / growth & development">
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Search in PubMed
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</a></li><li><a class="search-in-mesh-link dropdown-block-link"
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href="https://www.ncbi.nlm.nih.gov/mesh?term=Zebrafish"
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data-ga-category="search_mesh"
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data-ga-action="mesh_term_link"
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data-ga-label="Zebrafish* / growth & development">
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Search in MeSH
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</a></li><li><a class="add-to-search-link dropdown-block-link"
|
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ref="linksrc=add_to_search_link"
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role="button"
|
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data-search-term=""Zebrafish/growth and development"[MAJR]"
|
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data-ga-category="search"
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data-ga-action="add_to_search"
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data-ga-label=""Zebrafish/growth and development"[MAJR]"
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href="#">
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Add to Search
|
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</a></li></ul></div></div></div></li></ul>
|
||
</div>
|
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|
||
|
||
|
||
|
||
|
||
<div id="substances" class="substances keywords-section">
|
||
<h2 class="title">
|
||
Substances
|
||
</h2>
|
||
|
||
<ul class="keywords-list"><li><div class="keyword-actions dropdown-block"><button class="keyword-actions-trigger trigger keyword-link"
|
||
aria-label="Toggle dropdown menu for keyword Chromatin"
|
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tabindex="0"
|
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aria-controls="keyword-actions-substances-0"
|
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aria-expanded="false"
|
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data-pinger-ignore>
|
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Chromatin
|
||
</button><div id="keyword-actions-substances-0"
|
||
class="keyword-actions-dropdown dropdown dropdown-container"
|
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hidden
|
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aria-label="Dropdown menu for keyword Chromatin"><div class="title">Actions</div><div class="content"><ul class="keyword-actions-links"><li><a class="search-in-pubmed-link dropdown-block-link"
|
||
href="/?term=%22Chromatin%22%5Bnm%5D&sort=date&sort_order=desc"
|
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data-href="/?term=%22Chromatin%22%5Bnm%5D&sort=date&sort_order=desc"
|
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ref="linksrc=search_in_pubmed_keyword_link&keyword="Chromatin"[nm]"
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data-ga-category="search"
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data-ga-action="substance"
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data-ga-label="Chromatin">
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Search in PubMed
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</a></li><li><a class="search-in-mesh-link dropdown-block-link"
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ref="linksrc=search_in_mesh_link"
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href="https://www.ncbi.nlm.nih.gov/mesh?term=%22Chromatin%22"
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data-ga-category="search_mesh"
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data-ga-action="substance_link"
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data-ga-label="Chromatin">
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Search in MeSH
|
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</a></li><li><a class="add-to-search-link dropdown-block-link"
|
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ref="linksrc=add_to_search_link"
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role="button"
|
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data-search-term=""Chromatin"[nm]"
|
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data-ga-category="search"
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data-ga-action="add_to_search"
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data-ga-label=""Chromatin"[nm]"
|
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href="#">
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Add to Search
|
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</a></li></ul></div></div></div></li><li><div class="keyword-actions dropdown-block"><button class="keyword-actions-trigger trigger keyword-link"
|
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aria-label="Toggle dropdown menu for keyword Histones"
|
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tabindex="0"
|
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aria-controls="keyword-actions-substances-1"
|
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aria-expanded="false"
|
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data-pinger-ignore>
|
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Histones
|
||
</button><div id="keyword-actions-substances-1"
|
||
class="keyword-actions-dropdown dropdown dropdown-container"
|
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hidden
|
||
aria-label="Dropdown menu for keyword Histones"><div class="title">Actions</div><div class="content"><ul class="keyword-actions-links"><li><a class="search-in-pubmed-link dropdown-block-link"
|
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href="/?term=%22Histones%22%5Bnm%5D&sort=date&sort_order=desc"
|
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data-href="/?term=%22Histones%22%5Bnm%5D&sort=date&sort_order=desc"
|
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ref="linksrc=search_in_pubmed_keyword_link&keyword="Histones"[nm]"
|
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data-ga-category="search"
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data-ga-action="substance"
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data-ga-label="Histones">
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Search in PubMed
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</a></li><li><a class="search-in-mesh-link dropdown-block-link"
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ref="linksrc=search_in_mesh_link"
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href="https://www.ncbi.nlm.nih.gov/mesh?term=%22Histones%22"
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data-ga-category="search_mesh"
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data-ga-action="substance_link"
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data-ga-label="Histones">
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Search in MeSH
|
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</a></li><li><a class="add-to-search-link dropdown-block-link"
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ref="linksrc=add_to_search_link"
|
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